Research ArticleConfirmation, Sensory and Motor Systems

Layer-specific Developmental Changes in Excitation and Inhibition in Rat Primary Visual Cortex

Roberta Tatti, Olivia K. Swanson, Melinda S. E. Lee and Arianna Maffei
eNeuro 4 December 2017, 4 (6) ENEURO.0402-17.2017; https://doi.org/10.1523/ENEURO.0402-17.2017

Article Figures & Data

Figures

  • Figure 1.
    Figure 1.

    Age-dependent expression of interneurons in rat primary visual cortex. A, Confocal image taken with a 01× objective showing the location of the monocular portion of rat V1 (V1m). The landmarks of the white matter were used to identify V1m. Scale bar: 500 µm. B, Adjacent sections were processed with antibody staining for the vesicular glutamate transporter-2 (VGLUT2, green) and fluorescent Nissl (blue) to measure L4 distance from the pia and identify layers. Scale bar: 400 µm. C, High-magnification image acquired with a 20× objective showing PV+ (upper panel; magenta) and SST+ (SST; middle panel; green) expressing neurons; the merged image is shown in the lower panel. Scale bar: 35 µm. D, From left to right: confocal images showing the distribution of Nissl, PV+, and SST+ neurons at P14 across the cortical mantle of rat visual cortex. Scale bar: 125 µm. E, From left to right: confocal images showing the distribution of Nissl, PV+, and SST+ neurons at P17. Scale bar: 125 µm. F, Percentage of PV+ interneurons across V1 cortical layers quantified in four postnatal age groups (P14, P17, P21, P30). G, Percentage of SST+ interneurons across V1 cortical layers quantified in four postnatal age groups. 10 coronal slices from 4 rats were used for quantification. Data are mean ± SEM. Statistical significance was calculated using ANOVA, and p values were corrected for multiple comparisons.

  • Figure 2.
    Figure 2.

    Laminar differences in excitatory and inhibitory synaptic drive. A, From top to bottom: confocal images showing pyramidal neurons located respectively in L2/3, L4, L5, and L6. Yellow arrowheads indicate recorded neurons within the corresponding layer. It should be noted that multiple neurons were simultaneously recorded from different layers. Scale bars: 40 µm. B, After confocal imaging, slices were unmounted and developed with DAB to reconstruct recorded neurons using the software Neurolucida on a bright-field microscope. We included in the analysis only data obtained from neurons with pyramidal cell morphology and intact apical dendrite. Scale bar: 100 µm. C, From top to bottom: (upper panel, blue traces) representative traces of sEPSC recorded in L2/3, L4, L5, and L6 at P14 and P30; (lower panel, blue bar plots) average sEPSC charge quantified at P14, P17, P21, and P30. D, From top to bottom: (upper panel, red traces) representative traces of sIPSC recorded in L2/3, L4, L5, and L6 at P14 and P30; (lower panel, red bar plots) average sIPSC charge quantified at P14, P17, P21, and P30. E, Average excitatory/inhibitory (E/I ratio) ratio across layers in four postnatal age groups. Data are presented as mean ± standard error; asterisks indicate significant differences. Experimental values and statistics are reported in Tables 1 and 2.

  • Figure 3.
    Figure 3.

    Laminar differences in synaptic charge and E/I balance. A, Developmental time course of excitatory charge in L2/3 (blue), L4 (red), L5 (purple), and L6 (green). B, Time course of inhibitory charge in the different layers. C, Time course of the E/I ratio of the charges compared across layers. Statistical analysis for these data are provided in Table 3.

  • Figure 4.
    Figure 4.

    Maturation of spontaneous excitatory and inhibitory inputs in V1. A, Average input resistance (Rin) measured from V1m pyramidal neurons in L2/3, L4, L5, and L6 and in the different age groups (P14, P17, P21, P30). B, From left to right: average sEPSC frequency measured from pyramidal neurons grouped by layer and age group. C, From left to right: cumulative distributions of sEPSC amplitude from L2/3, L4, L5, and L6 pyramidal neurons in the four age groups. Small bar plots represent average sEPSC amplitudes. D, From left to right: average sIPSC frequency measured by layer and age group. E, From left to right: cumulative distributions of sIPSC amplitude from L2/3, L4, L5, and L6 pyramidal neurons in the four age groups. Small bar plots represent average sIPSC amplitudes. 100 events/neuron were included in the cumulative plots. Data are mean ± SEM. Values and statistics are reported in Tables 1 and 2.

  • Figure 5.
    Figure 5.

    Laminar differences in excitatory and inhibitory synaptic transmission. A, B, Time course of sEPSC frequency (A) and amplitude (B) compared across layers. L2/3: blue; L4: red; L5: purple; L6: green. C, D, sIPSC frequency (C) and amplitude (D) plotted by layer and age group. The statistical analyses for the data are reported in Table 3.

  • Figure 6.
    Figure 6.

    bDEO does not affect PV+ and SST+ neurons number in rat V1m. A, Diagram of binocular delayed eye opening (bDEO): both eyes were sutured shut before eye opening and kept closed until P17. B, From left to right: confocal images taken from control rats at P17 (Control) showing Nissl staining (blue), PV+ (magenta), and SST+ (green) immunostaining and merge. Scale bar: 125 µm. C, From left to right: confocal images taken from P17 bDEO rats showing Nissl staining (blue), PV+ (magenta), and SST+ (green) immunostaining and merge. Scale bar: 125 µm. D, Quantification of the percentage of PV+ interneurons in P17 control and P17 bDEO rats. E, Quantification of the SST+ interneurons from P17 control and P17 bDEO rats. 10 coronal slices from 4 rats were used to quantify PV and SST expression. Data are mean ± SEM.

  • Figure 7.
    Figure 7.

    Effect of 3- and 7-d bDEO on synaptic transmission and E/I ratio. A, Effect of bDEO on sEPSC charge, frequency, and amplitude plotted by layer and as percentage of control (represented by the dotted line). L2/3: blue; L4: red; L5: purple; L6: green. B, Laminar changes in sIPSC charge, frequency, and amplitude induced by 3- or 7-d bDEO. C, Laminar ratio of excitatory and inhibitory charge after 3- and 7-d bDEO. The statistical analyses for these data are reported in Tables 4 and 5.

  • Figure 8.
    Figure 8.

    The number and distribution of PV+ and SST+ neurons is not altered by mDEO. A, Diagram of mDEO: one eye was sutured shut right before eye opening (P13–P14) and kept closed until P21. B, From left to right: confocal images taken from the control hemisphere (ipsilateral to the closed eye) showing Nissl (blue), PV+ (magenta), and SST+ (green) staining and merge. Scale bar: 125 µm. C, From left to right: confocal images taken from the hemisphere contralateral to the closed eye (deprived) showing Nissl, PV+, and SST+ staining and merge. Scale bar: 125 µm. D, Quantification of the percentage of PV+ interneurons in control and deprived hemispheres across V1 cortical layers (data obtained from 10 coronal slices from 4 rats). E, Quantification of the percentage of SST+ interneurons in control and deprived hemispheres across V1 cortical layers (data were collected from 10 coronal slices from 4 rats). Data are mean ± SEM.

  • Figure 9.
    Figure 9.

    Effect of 3- and 7-d mDEO on synaptic transmission and E/I ratio. A, Effect of mDEO on sEPSC charge, frequency, and amplitude plotted by layer and as percentage of control (represented by the dotted line). L2/3: blue; L4: red; L5: purple; L6: green. B, Laminar changes in sIPSC charge, frequency, and amplitude induced by 3- or 7-d mDEO. C, Laminar ratio of excitatory and inhibitory charge after 3- or 7-d mDEO. The statistical analyses for these data are reported in Tables 6 and 7.

Tables

  • Table 1.

    Developmental changes in excitatory charge, sEPSC frequency, and sEPSC amplitude.

    A, Multiple comparisons across ages and KW-ANOVA.

    LayerCharge (nC)Frequency (Hz)Amplitude (pA)
    P14P17P21P30KW-ANOVAP14P17P21P30KW-ANOVAP14P17P21P30KW-ANOVA
    L2/35.8 ± 0.810.4 ± 1.95.6 ± 1.57.4 ± 1.1p = 0.22.3 ± 0.23.8 ± 0.62.9 ± 0.64.3 ± 0.5*p = 0.00721.6 ± 1.222.2 ± 2.116.7 ± 1.816.7 ± 1.7*p = 0.02
    L49.4 ± 1.513.8 ± 2.16.3 ± 0.710.2 ± 1.1p = 0.063.6 ± 0.45.6 ± 0.73.3 ± 0.55.8 ± 0.5*p = 0.00521.2 ± 1.419.3 ± 1.218.2 ± 1.018.3 ± 1.7p = 0.2
    L56.4 ± 1.17.6 ± 1.15.4 ± 0.58.0 ± 1.5p = 0.42.5 ± 0.52.9 ± 0.42.9 ± 0.53.7 ± 0.5p = 0.321.1 ± 1.421.0 ± 1.819.7 ± 1.518.6 ± 1.6p = 0.7
    L62.9 ± 0.54.8 ± 1.03.6 ± 0.93.2 ± 0.5p = 0.31.2 ± 0.32.2 ± 0.51.7 ± 0.42.1 ± 0.3p = 0.319.9 ± 1.219.9 ± 1.120.0 ± 1.114.2 ± 0.7*p = 0.002

    • Results are expressed as mean ± SEM.

  • B, MW and KS tests.

    ComparisonMW testKS test
    L2/3L4 FrequencyL6 AmplitudeL2/3 AmplitudeL4 AmplitudeL5 AmplitudeL6 Amplitude
    FrequencyAmplitude
    P14 vs. P17p = 0.04p = 0.9p = 0.02p = 0.8p = 0.3*p < 0.001*p = 0.0014*p < 0.001
    P14 vs. P21p = 0.5p = 0.06p = 0.5p = 0.6*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P14 vs. P30*p < 0.001p = 0.02*p = 0.005*p = 0.002*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P17 vs. P30p = 0.4p = 0.03p = 0.7*p = 0.002*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P21 vs. P30p = 0.03p = 0.7*p = 0.004*p = 0.001*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P17 vs. P21p = 0.3p = 0.09p = 0.03p = 0.9p < 0.001*p < 0.001*p < 0.001*p = 0.010

    • *Statistically significant. Only p values <0.008 are considered significant owing to Bonferroni correction for multiple comparisons.

  • Table 2.

    Developmental changes in inhibitory charge, sIPSC frequency, and sIPSC amplitude across age.

    A, Multiple comparisons across ages and KW-ANOVA.

    LayerCharge (nC)Frequency (Hz)Amplitude (pA)
    P14P17P21P30KW-ANOVAP14P17P21P30KW-ANOVAP14P17P21P30KW-ANOVA
    L2/316.4 ± 2.422.3 ± 3.331.6 ± 9.340.7 ± 6.9*p = 0.0021.5 ± 0.22.7 ± 0.43.1 ± 0.54.9 ± 0.4*p < 0.00133.6 ± 1.529.8 ± 1.530.9 ± 3.927.0 ± 2.4p = 0.07
    L49.0 ± 1.727.5 ± 3.428.8 ± 2.452.3 ± 5.8*p < 0.0011.1 ± 0.23.4 ± 0.43.2 ± 0.45.6 ± 0.3*p < 0.00129.7 ± 1.931.2 ± 2.131.4 ± 2.037.0 ± 2.1p = 0.2
    L515.1 ± 2.829.3 ± 3.742.8 ± 5.848.3 ± 7.2*p < 0.0012.2 ± 0.53.4 ± 0.44.5 ± 0.55.1 ± 0.4*p = 0.00227.4 ± 129.6 ± 2.731.2 ± 2.532.4 ± 2.7p = 0.7
    L63.6 ± 0.612.5 ± 2.516.4 ± 0.731.7 ± 4.6*p < 0.0010.5 ± 0.11.7 ± 0.41.9 ± 0.33.5 ± 0.4*p < 0.00125.6 ± 2.029.2 ± 1.529.1 ± 1.730.3 ± 4.1p = 0.4

    • Results are expressed as Mean ± SEM.

  • B, MW and KS tests.

    ComparisonMW testKS test
    L2/3L4L5L6L2/3 AmplitudeL4 AmplitudeL5 AmplitudeL6 Amplitude
    ChargeFrequencyChargeFrequencyChargeFrequencyChargeFrequency
    P14 vs. P17p = 0.2p = 0.01*p < 0.001*p < 0.001p = 0.01p = 0.09*p < 0.001*p < 0.001*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P14 vs. P21p = 0.07p = 0.01*p < 0.001*p < 0.001*p < 0.001p = 0.01*p = 0.007*p < 0.001*p < 0.001p = 0.035*p < 0.001*p < 0.001
    P14 vs. P30*p < 0.001*p < 0.001*p < 0.001*p < 0.001*p = 0.001*p = 0.002*p < 0.001*p < 0.001*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P17 vs. P30p = 0.01*p = 0.001*p < 0.001*p < 0.001p = 0.04p = 0.01*p = 0.002*p = 0.002*p < 0.001*p < 0.001*p < 0.001*p < 0.001
    P21 vs. P30p = 0.1p = 0.02*p = 0.001*p < 0.001p = 0.6p = 0.3p = 0.01p = 0.02p = 0.058*p < 0.001*p < 0.001*p < 0.001
    P17 vs. P21p = 0.5p = 0.6p = 0.9p = 0.8p = 0.08p = 0.1p = 0.3p = 0.4*p < 0.001*p = 0.002*p < 0.001p = 0.3

    • *Statistically significant. Only p values <0.008 are considered significant due to Bonferroni correction for multiple comparisons.

  • Table 3.

    Multicomparisons of excitatory and inhibitory synaptic transmission across layers.

    ComparisonsEPSC chargesEPSC frequencysEPSC amplitudesIPSC chargesIPSC frequencysIPSC amplitudeE/I ratio
    P14
        KW-ANOVA*p < 0.001*p < 0.001p = 0.8*p < 0.001*p < 0.001*p = 0.007*p < 0.001
        M-W U
            L2/3 vs. L4p = 0.03*p = 0.007p = 0.01p = 0.07p = 0.1*p < 0.001
            L2/3 vs. L5p = 0.7p = 0.9p = 0.6p = 0.4*p = 0.005p = 0.5
            L2/3 vs. L6p = 0.01p = 0.01*p < 0.001*p < 0.001*p = 0.004*p = 0.001
            L4 vs. L5p = 0.08p = 0.09p = 0.07p = 0.06p = 0.3p = 0.01
            L4 vs. L6*p < 0.001*p < 0.001*p = 0.005*p = 0.003p = 0.1p = 0.07
            L5 vs. L6p = 0.02p = 0.04*p < 0.001*p < 0.001p = 0.2p = 0.1
    P17
        KW-ANOVA*p = 0.01*p = 0.001p = 0.8*p = 0.001*p = 0.005p = 0.8p = 0.07
        M-W U
            L2/3 vs. L4p = 0.2p = 0.09p = 0.3p = 0.2
            L2/3 vs. L5p = 0.6p = 0.2p = 0.2p = 0.4
            L2/3 vs. L6p = 0.03p = 0.02p = 0.01p = 0.04
            L4 vs. L5p = 0.08*p = 0.005p = 0.8p = 0.7
            L4 vs. L6*p = 0.005*p < 0.001*p = 0.001*p = 0.002
            L5 vs. L6p = 0.1p = 0.1*p = 0.001*p = 0.003
    P21
        KW-ANOVAp = 0.1p = 0.2p = 0.5*p = 0.002*p = 0.009p = 0.8p = 0.4
        M-W U
            L2/3 vs. L4p = 0.6p = 0.6
            L2/3 vs. L5p = 0.1p = 0.08
            L2/3 vs. L6p = 0.1p = 0.3
            L4 vs. L5p = 0.1p = 0.08
            L4 vs. L6*p < 0.001p = 0.02
            L5 vs. L6*p < 0.001*p < 0.001
    P30
        KW-ANOVA*p < 0.001*p < 0.001p = 0.1p = 0.07*p = 0.01*p = 0.03p = 0.2
        M-W U
            L2/3 vs. L4p = 0.1p = 0.04p = 0.3*p = 0.004
            L2/3 vs. L5p = 0.8p = 0.5p = 0.7p = 0.1
            L2/3 vs. L6*p = 0.004*p = 0.001p = 0.03p = 0.8
            L4 vs. L5p = 0.2p = 0.01p = 0.7p = 0.1
            L4 vs. L6*p < 0.001*p < 0.001*p = 0.004p = 0.07
            L5 vs. L6*p = 0.007p = 0.02p = 0.02p = 0.4

    • Development: sEPSCs and sIPSCs; multiple comparisons across layers. *Statistically significant. Only p values <0.008 are considered significant due to Bonferroni correction for multiple comparisons.

  • Table 4.

    Changes in excitatory and inhibitory transmission after bDEO.

    LayerEPSC chargeEPSC frequencyEPSC amplitudeIPSC chargeIPSC frequencyIPSC amplitudeE/I ratio
    bDEOP17bDEOP21bDEOP17bDEOP21bDEOP17bDEOP21bDEOP17bDEOP21bDEOP17bDEOP21bDEOP17bDEOP21bDEOP17bDEOP21
    L2/351 ± 5; *p = 0.01126 ± 22; p = 0.564 ± 7; *p = 0.04107 ± 14; p = 0.888 ± 7; p = 0.3116 ± 10; p = 0.363 ± 9; *p = 0.0390 ± 13; p = 0.877 ± 7; p = 0.194 ± 12; p = 0.884 ± 5; *p = 0.04104 ± 7; p = 0.883.6 ± 12.1 p = 0.5134.8 ± 24.8 p = 0.3
    L460 ± 9; *p = 0.03128 ± 25; p = 0.357 ± 9; *p = 0.01118 ± 21; p = 0.6105 ± 8; p = 0.6106 ± 6; p = 0.364 ± 6; *p = 0.0199 ± 10; p = 0.970 ± 8; *p = 0.0299 ± 8; p = 0.994 ± 5; p = 0.5107 ± 2; p = 0.283.0 ± 9.5p = 0.4130.6 ± 23.4p = 0.3
    L559 ± 8; *p = 0.03122 ± 30; p = 0.563 ± 9; *p = 0.03107 ± 23; p = 0.895 ± 7; p = 0.799 ± 11; p = 0.959 ± 10; *p = 0.02101 ± 20; p = 0.956 ± 6; *p = 0.01105 ± 13; p = 0.896 ± 7; p = 0.799 ± 7; p = 0.9119.8 ± 26.9p = 0.5131.4 ± 43.8p = 0.5
    L682 ± 16; p = 0.5106 ± 18; p = 0.865 ± 11; p = 0.284 ± 21; p = 0.6111 ± 6; p = 0.2117 ± 10; p = 0.177 ± 12; p = 0.3114 ± 18; p = 0.160 ± 9; p = 0.0793 ± 14; p = 0.4114 ± 9; p = 0.1116 ± 8; p = 0.191.9 ± 13.5; p = 0.794.3 ± 9.6; p = 0.8

    • Results are expressed as mean % change from control (age-matched littermates) ± SEM. *Statistically significant (p < 0.05) by t test.

  • Table 5.

    Multiple comparisons of excitatory and inhibitory synaptic transmission after bDEO.

    bDEOsEPSC chargesEPSC frequencysEPSC amplitudesIPSC chargesIPSC frequencysIPSC amplitudeE/I ratio
    P17p = 0.8p = 0.9p = 0.06p = 0.7p = 0.2p = 0.08p = 0.8
    P21p = 0.9p = 0.6p = 0.5p = 0.6p = 0.8p = 0.6p = 0.7

    • Multiple comparisons across layers by KW-ANOVA. Values for bDEO P17 and P21 are reported in Table 4.

  • Table 6.

    Changes in excitatory and inhibitory synaptic transmission after mDEO expressed as % change from control neurons of the hemisphere contralateral to the open eye.

    LayersEPSC chargesEPSC frequencysEPSC amplitudesIPSC chargesIPSC frequencysIPSC amplitudeE/I ratio
    mDEO P17mDEO P21mDEO P17mDEO P21mDEO P17mDEO P21mDEO P17mDEO P21mDEO P17mDEO P21mDEO P17mDEO P21mDEO P17mDEO P21
    L2/350 ± 6; *p < 0.00173 ± 6; *p = 0.0358 ± 9; p = 0.00399 ± 6; p = 0.999 ± 10; p = 0.976 ± 5; *p = 0.01131 ± 23; p = 0.275 ± 7; *p = 0.02123 ± 16; p = 0.298 ± 4; p = 0.9108 ± 12; p = 0.683 ± 5;*p = 0.0440 ± 7; *p < 0.001104 ± 13; p = 0.7
    L4120 ± 9; p = 0.2164 ± 18; *p = 0.0197 ± 8; p = 0.897 ± 9; p = 0.9111 ± 7; p = 0.2130 ± 9; *p = 0.01147 ± 16; *p = 0.01147 ± 16; *p = 0.01101 ± 8; p = 0.9142 ± 16; *p = 0.04134 ± 8; *p = 0.003109 ± 7; p = 0.372 ± 9; *p = 0.04116 ± 14; p = 0.4
    L5132 ± 16; p = 0.1585 ± 16; p = 0.5131 ± 21; p = 0.372 ± 15; p = 0.588 ± 5; p = 0.193 ± 7; p = 0.580 ± 8; p = 0.270 ± 9; p = 0.0980 ± 8; p = 0.387 ± 7; p = 0.2103 ± 8; p = 0.776 ± 4; *p = 0.04160 ± 29; p = 0.1112 ± 15; p = 0.6
    L678 ± 16; p = 0.4177 ± 33; *p = 0.0271 ± 13; p = 0.2170 ± 19 *p = 0.00399 ± 7; p = 0.990 ± 5; p = 0.286 ± 18; p = 0.1697 ± 19; p = 0.980 ± 14; p = 0.5107 ± 14; p = 0.7106 ± 10; p = 0.6100 ± 8; p = 0.981 ± 21; p = 0.5171 ± 36; p = 0.06

    • Results are expressed as mean % change ± SEM. *Statistically significant (p < 0.05) by t test.

  • Table 7.

    Laminar specific effects in excitatory and inhibitory synaptic transmission following monocular delayed eye opening

    mDEOsEPSC chargesEPSC frequencysEPSC amplitudesIPSC chargesIPSC frequencysIPSC amplitudeE/I ratio
    P17
        KW-ANOVA*p < 0.001*p = 0.004p = 0.1*p = 0.009*p = 0.04p = 0.08p < 0.001
        M-W U
            L2/3 vs. L4*p < 0.001*p = 0.004p = 0.5p = 0.2*p < 0.001
            L2/3 vs. L5*p < 0.001*p = 0.003p = 0.1p = 0.06*p < 0.001
            L2/3 vs. L6p = 0.2p = 0.3p = 0.2p = 0.02p = 0.08
            L4 vs. L5p = 0.6p = 0.4*p = 0.001p = 0.1*p = 0.001
            L4 vs. L6p = 0.01p = 0.09p = 0.009p = 0.04p = 0.01
            L5 vs. L6p = 0.02p = 0.05p = 0.9p = 0.7p = 0.02
    P21
        KW-ANOVA*p < 0.001*p = 0.001*p < 0.001*p = 0.002*p = 0.04*p = 0.01p = 0.5
        M-W U
            L2/3 vs. L4*p < 0.001p = 0.7*p < 0.001*p < 0.001p = 0.03*p = 0.007
            L2/3 vs. L5p = 0.7p = 0.08p = 0.1p = 0.7p = 0.4p = 0.8
            L2/3 vs. L6*p = 0.001*p = 0.004p = 0.1p = 0.7p = 0.9p = 0.1
            L4 vs. L5*p = 0.003p = 0.09p = 0.008*p < 0.001*p = 0.003*p = 0.003
            L4 vs. L6p = 0.8*p = 0.006*p = 0.002p = 0.02p = 0.1p = 0.3
            L5 vs. L6*p = 0.003*p = 0.002p = 0.7p = 0.5p = 0.5p = 0.09

    • *Statistically significant. Only p values <0.008 are considered significant due to Bonferroni correction for multiple comparisons. Values for mDEO P17 and mDEOP21 are reported in Table 6.

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Layer-specific Developmental Changes in Excitation and Inhibition in Rat Primary Visual Cortex
Roberta Tatti, Olivia K. Swanson, Melinda S. E. Lee, Arianna Maffei
eNeuro 4 December 2017, 4 (6) ENEURO.0402-17.2017; DOI: 10.1523/ENEURO.0402-17.2017
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